Field Studies Council: Soil Biodiversity Course March 2015 Introduction to Collembola

Peter Shaw

Introduction

• Aim of the course: To introduce the main groups of UK

Collembola and to give you experience of working through

the FSC key to these animals, plus give you a chance to

collect a few specimens!

Resources: whatever Collembola we can collect, plus a

variety of specimens in alcohol and on slides that I have

accumulated.

Collembola (springtails) Taxonomic position

What this figures suggests is that a myriapod-

like ancestor evolved 6 legs, then one group

with muscular antennae gave the Collembola

diplura and protura.

Last I heard, this was OK except replace

‘ancestral myriapod’ with ‘ancestral

crustacean’!

All the zoology texts call these animals apterygota – wingless

insects. Hardly odd – they have 6 legs and run around on

land, but lack wings.

Recent DNA work has altered this; Collembola are probably closer to crustacea.

(When Peter Lawrence suggested this in 1996 the paper came with an editorial

effectively questioning his sanity!)

Collembola – crustaceans??

On the face of it, they’re not. Only 6 legs (on the thorax only), and just 2

antennae?

But their jumping organ (furcula) is fused abdominal legs, and latches onto the

tenaculum which looks like it could be homologous with fused legs. The

ventral tube looks like it derived from legs encasing gills.

The antennae are single, but there are 2 more cephalic sensory organ, the Post

Antennal organs PAO, which looks like the remains of the antennule.

It is possible that the last

common ancestor of springtails

and true insecta had >6 legs, as

do most crustacea.

Orchesella

cincta with

8 legs, after

1 HOX

gene is

disabled.

Collembola (springtails)

These are among the oldest (evolutionarily) and least changed of all

terrestrial arthropod groups. Most books still call them apterygote insects,

close to protura and diplura. A better model is to see the entire group

‘Pancrustacea’ (= hexapods plus crustacea) as a folded sheet of paper;

insects up one side, crustacea up the the other, and Collembola up the

middle! They moult continuously throughout life.

The surface dwelling forms have an escape mechanism involving a unique

jumping organ the furca (sometimes furculum), apparently fused vestigial

legs that insert on abd. IV. This latches into a hook (the tenaculum) on abd

III, stores energy and releases it to propel the animal’s jump.

The diagnostic feature of the class is the ventral tube or collophore, which

gave the group its name (Collembola = sticky peg).

Antenna, with

4 segments.

This genus

(Tomocerus) is

odd in having a

big 3rd

segment.

Eyes; <=8. also a PAO

Head +

mouth

Anus abd6,

genital

orifice abd 5

manubrium

Furca: dens

mucro

Legs

TN

Metathorax

Mesothorax

Prothorax, highly reduced

in many forms.

VT

Abdomen

segments 1-6

(last 2-3 may

fuse)

thorax

segments 1-3

PAO = Post antennal organ

TN: Tenaculum (hook for furca)

VT: Ventral tube or collophore

Collembola

anatomy

Modified from

an original by

R Fox

Collembola following this morphology are the superfamilies

poduromorpha and entomobryomorpha.

Poduridae

Isotomidae

Entomobryomorpha

Here Tomocerus

‘*’ shows the

prothorax

*

*

*

The symphypleona (sminthurids) follow this basic

plan but have fused body segments and look very

weird. They do have the furca and ventral tube, and

are usually <1mm.

About the only Collembola pests are in this group; the

lucerne flea Sminthurus viridis, which sometimes

nibbles clover.

When sminthurids jump and land on their back, they protrude their ventral tube to

2* body length.

Its end is sticky; it adheres to the substrate, and as they pull it in they right

themselves.

Allacma purpurescens

© the bald eagle,south Dakota Picture of a Dicyrtomina laying on its back using its everted sacs of

the VT to pull it back on its feet

© Jim McClarin, USA:

Here we see one of the 2 sacs of the ventral tube: these can be used to groom the

animal, to drink up water, and to pull itself up, to anchor: think of an elephant’s

trunk! The tube can be retracted quickly – a lucky photo. In alcohol these animals

look disembowelled, but aren’t!

Allacma fusca

frontal aspect, showing the ventral

tube or collophore and furca,

Hall, K. © 2005.

Sminthuridae sp.: collophore with eversed vesicles, ventral aspect.

SEM by Zeppelini, D. (2005).

Dicyrtomina saundersii

drinking using its collophore

Respiration – mainly cuticular, though some sminthurids have a

simple tracheal system. Their thin permeable cuticle makes them

creatures of damp habitats.

Reproduction: Males deposit a spermatophore, and females take it

into their genital aperture. Clearly this gives the female

considerable leeway in her choice!

It may also explain the complex mating dances that have evolved in

a few species, such as Deuterosminthurus pallipes. Sminthurides

males have specially modified antennae to hold the female during

mating.

Many springtails are partially or fully parthenogenetic, probably due

to systemic infection with Wolbachia.

Taxonomic overview

Class Collembola

Arthropleona (debateably taxonomically

valid)

(“normal springtails”)

Neelipleona

(few and tiny spp

but common)

Symphypleona Entomobryomorpha Poduromorpha

Class level

Order level

The oldest hexapods in the world Rhynie chert is one of the most famous (and inaccessible)

fossil deposits in the earth’s history. It was laid down in the

Devonian, c. 410MYBP, at a time when the most advanced

vertebrate was a fish, plants had just started to appear on

land, and top predators were eurypterids. (CO2 was 10*

modern levels too).

500MYBP 400 300 200 100 0=Now

Dinosaurs Carboniferous

Humans (+other

mammals) split from

reptiles around here

Rhynie Chert

Just to emphasise how the world has

changed, England was then still

separated from Scotland by a deep sea,

and both were in the southern

hemisphere, about the latitude of

modern New Guinea.

hominids

Timeline

Rhyniognatha hirsti

Id as true insect by

2 hinges in its jaw

Rhyniella praecursor

(lacks a protergite – isotomidae).

Scourfield’s

diagram (1940)

reconstruction

The Rhynie chert contains Springtail

fossils, plus a possible true insect.

Cretaceous - Miocene

We have many specimens of ancient springtails, almost wholly

due to the preservative properties of amber.

There are especially big deposits in the Dominican republic

(cretaceous) and the baltic (miocene and more recent), with many

species presumed to be climbing trees.

The ‘recent’ deposits have familiar genera (mainly Lepidocyrtus

and Seira), while cretaceous amber has new genera in new

families. Another K/T biodiversity loss?

Grinnellia ventis from amber from the Dominican

Republic, original and reconstruction 2006

© Christiansen, K. & Nascimbene, P.

How many species?

Globally? No idea! Actually 6500 has been

mentioned, but let’s aim lower.

How many in the UK? As recorder, my list has 392

species (ignoring synonyms).

Going down this list and removing fictional, dubious

and otherwise shaky records: 381 species.

Do I believe this? No! New spp are arriving, old ones

being split. In the Onychiurids we REALLY don’t know

about spp richness at all; the field was plagued with false

splits, but may hide molecular species.

19

Pronunciation There are no agreed standards for pronunciation! I’ve heard an US citizen talk

about Sye-loss-obby for what I call Sigh-Low-sigh-bee.

Dicyrtoma Dye – sir – tomer?

Dick – row – tomer?

So I put out a global

email asking how to

say this:

Dee – churr – tome –er

Dye – sir – tomer – a

[A as in “jam”]

Dee-zeer-to-mah

Notable, misc

• Some species form swarms, most commonly on snow (yes, even in UK – first recorded 1867). Achorutes covered a chalk wall in Mickleham with a swarm millions strong. 6 widely different spp have swarmed in the UK

20

Ceratophysella

swarm

A swarm of Ceratophysella bengtsonni Bedfordshire May 2013

21

Some neanurids are very colourful

Neanura takoensis from China

after www.yellowman.cn., 2007 ©

Tetrodontophora bielanensis

The biggest Collembola known is Tetrodontophora

bielanensis, from the Urals, peaking at 9mm long.

Recently some unusual defensive chemicals

have been identified in the giant springtail

Tetrodontophora bielanensis, which double as

alarm pheromones.

A common UK species, Neanura muscorum, secretes

1,3 dimethoxybenzene, phenol and 2aminophenol!

Pheromones

Much of these animals’ behaviour is linked to pheromones,

eg Hypogastruras have been shown to synchronise their

moulting by means of a pheromone. (They also tend to be

smelly in culture – some protective/pheromonal

compounds?).

They have aggregation pheromones, leading to multi-species

aggregations and terribly noisy ecological data! (It also

attracts predatory mites).

Basic modern Collembola body forms

I’m not going to try to guess which of these is ancestral, but

there are some clearly defined body forms now to be

encountered:

Surface dwelling entomobryomorpha:

medium-large, coloured, almost

invariably with furcula (often well

developed)

Surface dwelling poduromorpha:

medium-large, coloured, generally

with furcula (but often small).

Surface dwelling symphypleona

medium-tiny, coloured, invariably with

well developed furcula.

Euedaphic poduromorpha: white,

eyes reduced or absent, furcula

tiny or (generally) absent.

Euedaphic symphypleona :

white, reduced eyes, furcula

present.

Basic Collembola body forms

Surface dwelling Entomobryomorpha:

These are often large (>2mm) forms with distinct colour patterns

and well developed jumping organ. Commonly encountered

genera are Entomobrya, Lepidocyrtus, Tomocerus, Isotoma.

Entomobrya Lepidocyrtus cyaneus

Tomocerus

Basic Collembola body forms

Surface dwelling poduromorpha: medium-large,

coloured, generally with furcula (but often small).

Less conspicuous than jumping forms but very

common and widespread – Neanura muscorum is

ubiquitous, Brachystomella in disturbed sites,

Xenylla up surfaces

Brachystomella

parvula

Lathyriopyga longiseta

Xenylla maritima

Neanura muscorum

3mm long

Basic Collembola body forms

Surface dwelling symphylopleona medium-tiny, coloured, invariably

with well developed furcula. “Lucerne fleas” (strictly = Sminthurus

viridis).

Sminthurus viridis

Sminthurinus trinotatus

Sminthurinus elegans

Dicyrtomina sp

Ptenothrix atra

Basic Collembola body forms

Euedaphic poduromorpha: white, eyes reduced or absent, furcula

tiny or (generally) absent. The hardest group to ID, probably not

valid until molecular systematics catch up with field-based reality!

Protaphorura (formerly

Onychiurus) armata

Mesaphorura (formerly

Tullbergia) sp

Basic Collembola body forms

Euedaphic symphylopleona : white, small-tiny, reduced eyes, furcula

present.

Neelides murinus Megalothorax

minimus (next to

Pogonognathellus

longicornis)

Lifestyles: feeding

Most Collembola are “detritivores”. This comes from seeing

them in leaf litter feeding on indeterminate particles, and finding

a variety of particles + soil microbes in their guts.

Actually, when given a choice springtails are quite

discriminating about food choices, with a preference for yeasts

and other non-toxic fungi. They will also culture well on the

green algae from tree bark (Pleurococcus).

Pests? There are few, but persistent, reports of springtails causing

agricultural damage. These are invariably a combination of high

numbers and animals attacking delicate young seedlings or leaves.

There have been a few cases of stems being girdled, but reduced leaf

area is commonest. At worst a nuisance.

Sminthurus viridis does explode to pest proportions in New Zealand

where it damages clover and alfalfa.

Humans?

Bizarrely there are a few cases in the wilder medical literature attributing

human parasitoses to springtails. These are actually examples of delusory

parasitoses – purely imagined, and the springtail-like shape in one flake of

skin an artefact of preparation. I have been asked to survey a client for this

once (until the university insurers heard the phrase “collect medical samples

from a psychiatric patient”). Twaddle, they’re imagining it. (I found one

paper about skin collembola that referred to their pupae!) Search on

Morgellons for a related delusional condition.

There is one case of an entomologist who pooted too hard and got Isotoma

olivacea eggs in his nose – they hatched, 50 emerged, and made him quite unwell

for a while.

The only credible Collembola-human zoonosis I’ve heard of! (One case of a

woman with a scalp infestation was traced to an entomobryid-laden plant pot by

the head of her bed!!)

http://www.bioone.org/action/showFullPopup?doi=10.1664%2F0028-7199%282004%29112%5B0087%3ACSAHEF%5D2.0.CO%3B2&id=_e2

http://www.usfreeads.com/538138-cls.html There are three main types of skin parasites that often get misdiagnosed by the medical community. If

you have felt like something was crawling on your skin or biting you but your doctor or dermatologist

said there is nothing wrong, it may just be that they don't know how to properly look for scabies,

collembola and morgellons disease.

SYMPTOMS OF A COLLEMBOLA (SPRINGTAIL) INFESTATION

Fatigue

Brain Fog, memory loss, difficulty thinking or concentrating

Mood Swings & Depression

Joint swelling and pain all over your body; Fibromyalgia

Visual decline

Autoimmune decreases

Hair Loss

Lesions that hurt and include inflammation

Hard Nodules under the skin

Itching, creepy crawly feeling

Organisms biting, moving & scratching under the skin

Sores that do not heal

Sleep Disorders Over abundance of lint or dust in the house; believed to be moulting by the springtail.

Are scabies and springtails the same? No, they are different parasites altogether. There is a big difference on how

the scabies mite and the springtails move around. Springtails have more of a jumping action; where the scabies can

only move approximately 2.5 cm per minute. It is not uncommon for people to see a springtail, if the light is just right,

to jump from an arm to a leg.

Additionally, we believe the skin crawling sensation may be due to the multiple flagella on the organism and the fibers

are produced by the organism itself. One researcher claims that the organism is found widespread in bottled spring

water from France. One study revealed that 1/3 of bottled waters are contaminated with this organism.

Gold standard

unmitigated paranoia-

inducing lies!!

36

Old soil biology texts claimed soil animals to be very non-fussy,

on the basis of the wide variety of material found in their guts.

This involves a rather crude operation known as ‘gut-squashing’.

In fact if you offer most springtails a choice of foods, they

routinely turn out to be highly selective:

Fungus 1 Fungus 2

Control

(agar)

From some expts I ran

in my PhD

Fungi avoided

included ‘poison pie’

and fly agaric.

Coincidence?

One Collembolan, a small slow moving dumpy little things called

Friesea mirabilis, is apparently an active predator of soil nematodes

(I have no idea how).

Recent work has shown that a common (edible) toadstool, the

ectomycorrhizal symbiont Laccaria bicolor is able to kill and

digest soil animals and transfer the nitrogen back to the host tree.

(As a twist, mycelium of a closely related fungus is readily eaten

by many soil arthropods).

Habitats: soils

In terms of species richness, the majority of Collembola live

in the soil in or below the litter layer, declining rapidly as one

enters the mineral soil.

You would expect that (like plants and many invertebrates)

contrasting soil/litter types would have contrasting

Collembola communities. To an extent this is true, and forms

the basis of many small projects. Where the contrasts are

sharp (eg lagoon -> ash dump -> woodland) one can easily

show community development.

In fact this effect is rather weak, with only a minority of

species showing clear habitat associations. There are

soome, eg Tetracanthella wahlgreni is a glacial relic

confined to high cold sites, while Folsomides turn up in

quarries that bake dry.

I also recall in York under Mike Usher, I

was studying acid wet pine woodland in

the borders and Richard next to me looked

at a dry calcareous chalk quarry near York.

We had several species of Collembola in

common! One (Isotoma viridis) is now a

group of 3 species, but the point about low

specificity stands.

Tetracanthella wahlgreni

BenMacdui 2011, c. 1200m

By the sea

Unsurprisingly for a near-crustacean radiation, some

Collembola are marine. None swim, and only a few are

evolved to survive tidal immersion.

The best known of these is Anurida

maritima (once Lipura maritima), which

is probably ubiquitous on our rocky

shores.

These grey/black animals emerge at low tide to graze (about 1 hour

after the rocks start to dry, in my experience); at high tide they

retreat to a cavity where their water-repellent hairs act to trap air.

Anurida maritima

There are another 5 similar things below the

tideline (2 Anurida and 3 Anuridella) so it’s

worth grabbing a few on sight to check ID.

You could easily add dots to the UK map!

One springtail lives under sea water for long

periods – months, maybe life? Cryptopygus

clavatus (formerly Proisotoma

buddenbrocki) lives under stones in rock

pools and can be kept submerged all winter

happily grazing the biofilm.

Anurida

granaria

Kew bridge,

inter-tidal

zone

16xi2012

43

Seaweed can turn up other Collembola in other families, eg the

coastal ‘counterpart’ to Entomobrya nivalis is Mesentoma dollfusi,

which seems scarce but is probably present on every beach in the

south of England!

Mesentoma dollfusi

In fresh water!

• Actually very few live IN freshwater, but Agrenia bidenticulata is fairly common and confined to stones in acid upland streams.

Rather more spp live ON freshwater, famously Podura

aquatica but also the tiny Sminthurides (with a gutter

shaped mucro to catch the water film). This is the

Neustonic community. (The Neuston is the

community of animals associated with the surface film

of water).

Agrenia bidenticulata

(Folsomia sp behind)

Podura aquatica

Sminthurides aquaticus

Up trees! Rather bizarrely, springtails are incorrigible tree climbers! Not all species, but

canopy communities can have the odd soil species.

The commonest tree climbers are large scaly species in the genera Tomocerus

s.lat., Lepidocyrtus and Orchesella, plus Entomobrya (large and hairy though not

scaly) and Vertagopus (waxy).

Entomobrya

albocincta –

rarely found

in the soil.

Up plants Leaf surfaces are the preferred habitat of some sminthurids, notably Sminthurus viridis,

also Deuterosminthurus and Bourletiella species. These are often missed by soil core

collection and are tiny to see, but vacuum up well!

Steve Hopkin mentions that while writing his book

on Collembola, there was a gentle stream of

Deuterosminthurus pallipes falling out of the bushes

onto his laptop – notable also for their complex

mating dances.

Katiannidae sp nov

Richmond park,

April 2009

Deuterosminthurus pallipes

mating dance Bourletiella arvalis

In ant nests

Bizarrely there are a few Collembola that associate with ants.

The commonest UK myrmecophile is Cyphoderus albinus, a

large blind white animal fairly close to Entomobrya. We also

have a rare myrmecophilous Entomobryid (Entomobryoides

myrmecophilus).

When I checked a load of ants nest springtails what turned up

with C. albinus was Lepidocyrtus cyaneus (not supposed to!)

Cyphoderus albinus

In Caves

Troglobites – species only found in caves

Oligaphorura schoetti; Folsomia agrelli, Disarrhopalites patrizi

Disparrhopalites patrizi

(I am indebted to the UK expert on cave biota: Graham

Proudlove)

Schaefferia lindbergi plus juvenile Oligaphorura

schoetti top right © P Chapman

(in Ogof Ffynnon Ddu )

Oligaphorura

schoetti on raft of

exuviae

Troglophiles – also have viable populations above

ground, eg Heteromurus nitidus, Folsomia candida

Folsomia candida

Heteromurus nitidus

50

The deepest-dwelling land animal is a springtail Plutomurus

ortobalaganensis, 1980m deep in the world's deepest cave Krubera-

Voronja, near the Black Sea in Abkhazia (Georgia.) along with three

new species of Collembola: Anurida stereoodorata, Deuteraphorura

kruberaensis and Schaefferia profundissima.

(Image: Rafael

Jordana and

Enrique Baquero

Snow fleas A few species have anti-freeze compounds that allow them to survive well below

freezing; snow fleas (Hypogastrura harvei and nivicola) actively graze snow algae

on the surface of snow in mountains and the arctic in winter! Hypogastrura cocklei

is the golden snow flea.

[Do not confuse these “snow fleas” with the winter-active mecopteran with the

same name]

Hypogastrura spp or ‘snow fleas’ swarming on snow

Just in 2008 the amino acid used as an antifreeze was synthesised and found to

have no sequence similarity to any known protein. (This amino acid is being used

to inform edible antifreezes to put into ice cream).

A swarm of Ceratophysella sigillata in

late winter. (Part of an annual cycle

with changes in morphology called

epitoky)

In Europe the ‘snow flea’ has been said to be “Isotoma nivalis”, though this species appears to be fictional! Desoria saltans is a valid name for a winter-active isotomid, and Frans Jansenns has seen a snow swarm of Isotoma viridis. I have seen a report that in 1927 Ernst Handschin found people trying to extract food oil from snow-surface swarms of Hypogastrura longispina in the alps (better than starving, probably, just!)

Probably quite a few species will swarm on snow given correct conditions.

54

Luminous Collembola

It is little known but a few Collembola are among the truly

luminous life forms. This is best known for Anurida

granaria: Arne Fjellberg found one glowing in a riverbank

in Iceland. In New Zealand they may be collected and

cultured.

http://www.byteland.org/biolumi

nus/methodology.html

Lobella sp, Japan http://xa.yimg.com/kq/groups/3650837/564

405651/name/2011_The%20Terrestrial%20

Bioluminescent%20Animals%20of%20Japa

n.pdf

People behind this course

(generally far behind…)

Sir John Lubbock MP, naturalist

30/4/1834 – 28/5/1913

(later Baron Avebury)

As an MP he introduced

many banking reforms

including the introduction

of bank holidays.

As a naturalist he produced a

magnificent work on UK

Collembola, with engravings

specially commissioned from a

deaf & dumb man called Mr

Hollick

How doth the Banking Busy Bee

Improve his shining Hours?

By studying on Bank Holidays

Strange insects and Wild Flowers!

Richard Siddoway Bagnall

(14 July 1889, Winlaton near Whickham -19 January 1962)

Was a hotel inspector by trade, odd but meant he travelled around the UK,

collecting insects (especially apterygota) as he did. He also described

new species, and probably got many of them right (but his descriptions

were rather vague, leaving a question mark over many species that look

to be UK endemics.)

I have seen him called a dipterist, and also a specialist in the

thysanoptera. (One of his species has turned up as a pest in Brazil

recently).

Other names that pop up extensively in 20th Century UK

Collembola records/literature:

H. Gisin – Switzerland; a unified European key, still cited.

HE Goto

Peter Miles

HJ Gough

Peter Lawrence, NHM

Mike Usher, York

Steve Hopkin 1956 -19/5/2006

It should have been Steve giving this course; Steve

was a lecturer at Reading university with a wide

interest in invertebrates. He was killed in a car crash

in 2006 in Cornwall, where he was about to retire.

When I took over his files about Collembola

distribution (which remain definitive, though aging

slowly) I was approached by cornish spider experts

and the UK millipede group, both of whom had

found his work invaluable, which ecotoxicology

journals gave him an obituary all about his work on

pesticides and heavy metals.

Arne Fjellberg

61

Very much alive and kicking, Arne produced a

series of keys to Collembola in his native Norway,

with an interest in arctic and high-altitude species.

Arne has published many detailed taxonomic papers describing and

re-describing new and poorly-defined species of Collembola, and

gave Steve Hopkin much valuable information, especially when

unravelling some of Bagnall’s descriptions and relating the UK fauna

to the wider european fauna.

Frans Janssens

62

Frans is based in Antwerp, Belgium, and has become

involved in photo-ID of Collembola on the Flickr

photo-sharing website.

He runs the immensely useful webpage

www.Collembola.org

Featuring photos and online keys. He has also added

species to the UK list by online macro-photography

groups.

Matty Berg – Vrije University,

Amsterdam

Matty’s research has a focus on the explanation of soil fauna

community composition over time and across space. He

studies the importance of species traits (incl. variability and

plasticity) for species interactions. This involves, among

other experiments, adding various Collembola species by

pooter to microcosms, to explore their effects on community

composition and key ecological processes. He has build a

large trait databases for Collembola that helps him to

understand community changes.

He has also published a series of papers on unusual

springtails in the Netherlands, where he monitors and maps

the biogeography of Collembola and other soil invertebrates

(Isopods, Diplopoda, Chilopoda). He is currently the

coordinator of the Collembola survey group in the

Netherlands.

Myself

With a degree in zoology I undertook a PhD on Collembola of lodgepole

pine plantations with Mike Usher of York university. The aim was to see

whether they grazed the hyphae of ectomycorhizal fungi, thereby

damaging the trees. (Work based in Spadeadam forest, borders). That

turned out to be fearsomely tricky to answer and even now the answer is

along the lines of “probably, but not much”.

After submitting my PhD I started work with the CEGB on acid rain. Sadly I

never got the time or funding to link springtails and acid rain – blame NERC

for not funding it !!

Latterly I did acquire an interest in the biodiversity of ex-industrial sites (PFA),

and have found nice successional changes in Collembola corresponding to soil

development.

Roehampton!

In 1991 Thatcher privatised us, and in 1992 I started at Roehampton. The acid rain

work needed a team of technicians that had just blown apart, so I returned to

Collembola (inter alia). I collaborated somewhat with Steve Hopkin, realised what

a huge amount he’d done and how not to trust my IDs without double checking!

If we have a living link it’s my ex-PhD student Erica McAlister, now NHM – we

took some of her specimens to Steve to check ID.

Then he died unexpectedly in 2006 after writing the FSc Collembola key but before

it was published. I was able to proof check and correct the manuscript (virtually no

typos or silly mis-numberings; quite remarkably well put together). We rescued

enough files to put Steve’s card index onto the NBN gateway, and I am UK

recorder for Collembola. Each paper has had to be retyped into a database to cross

link records to sources!

What we’re going to do: You have some Tullgren extracts, and later we will see what

comes out.

Today – intro to the key and to some common species. If

weather allows, some D-Vac / beating arboreal collection

After lunch – further specimens, not all so easy.